Vertebrate Zoology 60 (2) 2010 123 123 – 138 © Museum für Tierkunde Dresden, ISSN 1864-5755, 15.09.2010

Description of Dicrossus foirni sp. n. and Dicrossus warzeli sp. n. (Teleostei: Perciformes: Cichlidae), two new from the Rio Negro and the Rio Tapajós, Amazon drainage, Brazil

UWE RÖMER 1, INGO J. HAHN 2 & PABLO M. VERGARA 3

1 University of Trier, Inst. Biogeography, Dep. Geo-Sciences, Am Wissenschaftspark 25 – 27, D-54296 Trier, Germany eu.roemer(at)t-online.de (corresponding author) 2 Biogeography and Ecology Res. Group, Institute of Landscape Ecology, University of Münster, Robert-Koch-Str. 26 – 28, D-48149 Münster, Germany 3 Department of Engineer Geography, Universidad de Santiago de Chile, Av. Lib. B. O’Higgins 3363, P.C.: 7254758, Santiago, Chile

Received on June 19, 2010, accepted on July 30, 2010. Published online at www.vertebrate-zoology.de on September 02, 2010.

> Abstract Dicrossus foirni sp. n. and Dicrossus warzeli sp. n. are described from the Brazilian Amazon River system. Both species are differentiated from all other Neotropical by exhibiting three rows of longitudinal lateral spots. Dicrossus foirni sp. n., which is distributed in the northern Rio Negro drainage, exhibits a pattern of lateral spots formed by the clustering of individual dots. By contrast, Dicrossus warzeli sp. n., which is endemic to the Rio Tapajós, exhibits three longitudinal rows of horizontal dash-like spots on the sides of the body. Furthermore, the two species are distinguished by having different relationships between body height and body length.

> Resumen Dicrossus foirni sp. n. y Dicrossus warzeli sp. n. son descritas en la region del Río Amazonas. Ambas especies se differencian de todos los otros ciclides neotropicales en la composición y localización de sus tres bandas laterales. Dicrossus foirni sp. n., que se distribuye en la región Norte del Río Negro, muestra un patrón de manchas laterales la cual es formada por grupos de puntos individuales. Al contrario, el Dicrossus warzeli sp. n, que es endemico del Río Tapajós, muestra bandas delgadas continuas a ambos lados del abdomen. Además, ambas especies se distinguen por tener diferentes relaciones entre la altura corporal y el largo corporal.

> Kurzfassung Dicrossus foirni sp. n. und Dicrossus warzeli sp. n. werden aus dem brasilianischen Amazonaseinzug beschrieben. Die beiden Arten unterscheiden sich durch eine Anordnung von jeweils drei länglichen Fleckenreihen von allen anderen neotropischen Cichliden. Dicrossus foirni sp. n., die im nördlichen Rio Negro-Einzugsgebiet verbreitet sind, sind durch aus Einzelfl ecken zusammengesetzten Lateralmuster gekennzeichnet. Die im Rio Tapajós endemischen Dicrossus warzeli sp. n tragen hingegen Reihen durchgehender schmaler Längsstriche auf den Körperseiten. Beide Arten sind außerdem durch im Vergleich zur Kopfbreite unterschiedliche relative Körperhöhe voneinander zu unterscheiden.

> Key words Systematics, , biogeography, freshwater ecology, Neotropics, Amazon River, new taxa. 124 RÖMER et al.: Description of two new cichlid species from Brazil

Introduction Stawikowski & Werner (2004), and Staeck (2003). In this work only a few illustrations and notes on some basic colour patterns useful for identifi cation will be The Neotropical cichlid genus Dicrossus Steindach- included. GPS data for station F9/91R were taken with ner, 1875 at present includes three described dwarf a Garmin GPS 50 (Garmin Int. Inc., Lenexa, USA), and cichlid species: Dicrossus maculatus Steindachner, extracted from ONC Flight Navigation Charts (Ministry 1875 (the type species of the genus), Dicrossus fi lamen- of Defence, UK) and Google Earth for the igarapé near tosus (Ladiges, 1958), and Dicrossus gladicauda Schin- Sao Luiz. Statistical analysis was carried out using the dler & Staeck, 2008. Two more species, both from the PC program Statistika 6.0 for Windows (StatSoft Inc. Brazilian Amazon basin, have been known for at least Tulsa, USA). two decades (Kullander, 1990; Römer, 2000, 2006; Stawikowski & Werner, 2004). In this paper these two species, known since 1981 under various – in some Comparative material cases confusing – provisional names [i. e. Dicrossus sp. “A” (Kullander, 1990) Dicrossus sp. “Peru” (Tomey, Crenicara punctulata: MTD F 32159, 1 male, 81.8 mm SL, 1 1983), Dicrossus sp. “Rotfl ossen” (Linke & Staeck, female, 60.1 mm SL, collected March 1994 by M. Wöhler, 1992), Dicrossus sp. “Rio Negro” (Windisch, 1992), Anavillhanas archipelago, lower Rio Negro, federal state of Dicrossus sp. “Doppelfl eck” (Mayland & Bork, 1997), Amazonas, Brazil. and Dicrossus sp. “B” (Kullander, 1990), Dicrossus Crenicara latruncularium: UR.2010.06.113, 1 male, 1 female, sp. “Tapajós” (Warzel, 1996), Dicrossus sp. “Tapaios” not measured, commercial import, September 2009, C. Eon, (Warzel, 1996)], will be formally described below. Bordeaux. Dicrossus fi lamentosus: MTD F 32160, 6 specimens, 27.9 mm SL to 37.4 mm SL, collected October 13th 1991 by A. Schneider. Location: few hundred meters upriver from the mouth of the Iga- Material and Methods rapé Prósperitáte (station F9/91R: 00°09′47′′S / 64°07′11′′W), a small right hand tributary of the Rio Preto near Lago Urubu about 12.5 km NNW from the junction with the Rio Negro, federal Measurements and counts were taken as described state Amazonas, Brazil. 35 specimens (UR 2010.Df.1001) from in detail in Römer (2006) and Römer et al. (2003, vicinity of Puerto Inirida, imported by Aquapool (Wuppertal), 2004, 2006) All type specimens were preserved in 75% March 2007. ethanol. A small number of fi sh were preserved in the Dicrossus gladicauda: 80 specimens: MTD F 32155, 13 speci- fi eld, the rest after various periods of maintenance in mens, 22.8 mm SL to 37.7 mm SL; CAS 229 773, 13 specimens, the aquarium. The latter specimens were treated before 26.0 mm SL to 39.5 mm SL; CAS 229 774, 15 specimens, 23.7 preservation as described in detail by Römer & Hahn mm SL to 38.9 mm SL; MTD F 32156, 10 specimens all male, (2008). Voucher specimens were deposited in the fi sh 24.9 mm SL to 42.1 mm SL; MTD F 32157, 14 specimens, collections of the Museum for Natural History, Dresden 31.2 mm SL to 43.0 mm SL; MTD F 32158, 5 males, 36.4 mm (MTD F), California Academy of Sciences, San Fran- SL to 41.5 mm SL, cleared and stained; UR.2010.06.112, 10 cisco (CAS), Museu de Zoologia da Universidade de specimens, 26.0 mm SL to 40.6 mm SL. All from Rio Vichade, São Paulo (MZUSP), and the personal collection of the Eastern Colombia, collected May 2002. senior author (UR). Tissue samples for future genetic Dicrossus maculatus: UR.2010.06.113, 1male, 1 female, not analysis have been taken from all specimens, preserved measured, commercial import, October 2005. in pure ethanol, and registered under the same individual Supplementary material (non types). 6 live specimens (not numbers as the originating specimens. Descriptions are preserved) of each of both Dicrossus species described below, based mainly on holotypes, with additional informa- maintained in the aquarium for behavioural and ontogenetic tion taken from all paratypes. Information for colour observation, to be kept in the personal collection of UR after descriptions of voucher specimens was taken from fi sh preservation, and fi gures in Römer (2000, 2006). preserved in 75 % ethanol. Colour of dry specimens in some cases differs signifi cantly. Römer (2000, 2006) has presented numerous photographs of the species described herein. These fi gures are included as sup- Dicrossus foirni sp. n. plementary pictorial reference material. Behaviour was observed in the laboratory under conditions given in Figs. 1 – 7, 17 & 18, Tabs. 1 – 3) Römer & Hahn (2008). The coloration of live specimens is well known. It may be seen from numerous colour photographs in Holotype. (fi g. 1) MZUSP 106589, female, 42.8 mm SL, col- various publications, especially Römer (2000, 2006), lected by A. Schneider, October 15th 1992. Location: few hun- Vertebrate Zoology ■ 60 (2) 2010 125

Fig. 1. Dicrossus foirni sp. n., holotype (MZUSP 106589), female, colour pattern in 75 % ethanol, about six months after preservation.

Fig. 2. Dicrossus foirni sp. n., topotype (MTD F 32146), male, colour pattern in 75 % ethanol, about six months after preservation.

dred meters above mouth of the Igarapé Prósperitáte (station Diagnosis. Dicrossus foirni sp. n. is a medium-sized F9/91R: 00°09′47′′S / 64°07′11′′W), a small right-hand tributary crenicaratine cichlid species (males up to 71 mm SL, of the Rio Preto near Lago Urubu about 12.5 km NNW from females to 45 mm SL) sharing all typical characteristics the junction with the Rio Negro, federal state of Amazonas, of the genus Dicrossus Steindachner, 1875 as stated in Brazil. particular by Kullander (1990). The species exhibits Paratypes. 6 specimens. MTD F 32146 (topotype), male, pronounced sexual size and colour dimorphism. It is 70.9 mm SL (fi g. 2), and MZUSP 106590, male, 57.9 mm SL, unmistakeably characterised by three rows of round to M. Geismann, S. Leissner, and A. Schneider, other data as for vertical-ovoid, alternating dark grey to blackish, double holotype. MTD F female, 32147, 35.0 mm SL, MTD F 32148, or triple spots on the sides of the body; adult males with sex undetermined, 26.0 mm SL, cleared & stained, CAS 220 a black spot on the dorsum and basal part of the dorsal 433, male, 45.8 mm SL, CAS 220 434, female, 37.5 mm SL, fi n at the position of hard rays 7 to 8, and a pointed collected by Martin Wöhler, March 9th 1994, in a shallow swamp rhombic caudal fi n. lake in the Igarapé Prósperitáte about 1 kilometre upriver from its junction with the Rio Preto, about 1 hour upstream from the junction with the Rio Negro, federal state of Amazonas, Brazil. 126 RÖMER et al.: Description of two new cichlid species from Brazil

Fig. 3. Dicrossus foirni sp. n., adult male, dominant, territorial, not preserved.

Fig. 4. Dicrossus foirni sp. n., adult female, dominant, in brood care coloration, not preserved.

Description. Morphological data taken from 7 speci- in adult males slightly concave vertically below orbit. mens (26.0 to 70.9 mm SL). For biometric data see Abdominal contour almost straight in small individuals, tables 1 & 2, for meristic data see table 3. slightly to progressively convex in medium-size to large specimens of both sexes. Head elongate (about 31 % Habitus. (fi gs. 1 – 7, 17 & 18) Body moderately elon- of SL), in adults comparatively deep (up to 26.5 % of gate, laterally compressed, about two to two and a half SL), with (depending on SL) small subterminal, nar- times deeper than wide, signifi cantly deeper in adult row, and pointed mouth (fi g. 5). Positive allometry of males and females than in small to medium-size speci- snout length (about 5.2 % of SL in small specimens up mens (depth 34.1 % of SL vs. up to about 37 % in to nearly 11 % in large). Premaxillary fold of lip very adults of both sexes). Dorsal contour from tip of snout short. Distal edge of preoperculum serrated, with 33 to to posterior end of dorsal-fi n base regularly arched in 54 denticuli. Cheeks completely scaled (fi g.6). adult males, in smaller specimens and females only pre- Dorsal fi n [D. XV.7 (n = 6), XVI.6 (n = 1)] about half dorsally curved, in larger males interrupted by concave as high as body in juveniles and females to about one interorbital notch. Preventral contour generally straight, third of depth of body in adult males. Membranes not Vertebrate Zoology ■ 60 (2) 2010 127

extending above fi n spines, tips rectangular. Dorsal-fi n spines increasing rapidly from fi rst to seventh, which more than three times longer than fi rst; spines seven to ten of about equal length, posterior spines about 10 % longer than spine ten, except last which even longer. Soft portion of dorsal fi n pointed in males, rounded in females. Caudal fi n [C. 16 (n = 7)] in adult males rhombic and pointed, in medium-size males spatulate to rhombic, and rounded in smaller males and in females. Anal fi n (A: III.6 (n = 1), III.7 (n = 6) pointed in males, rounded in juveniles and females. Ventral fi ns [V. I.5 (n = 7)] short and rounded in juveniles and females (20 to 26 % of SL), extending back to genital papilla; in large males pointed with long fi lamentous extensions Fig. 5. Dicrossus foirni sp. n., juvenile male, frontal view of (up to 56 % of SL), extending back to soft part of anal- head showing the small terminal, narrow, and pointed mouth, fi n base. Pectoral fi ns [V. 11 (n = 6), 12 (n = 1)] rounded not preserved. in all specimens examined. Scales in E1 row 24 (n = 2), 25 (n = 3), or 26 (n = 2). Upper lateral line (14 to 18 scales) continu- ously arched from head to beginning of soft portion of dorsal fi n. Lower lateral line irregular (3 to 10 scales), in some cases interrupted, extending straight to base of caudal fi n. Teeth in three series in both jaws; in large speci- mens, especially males, series across anterior part of lower jaw irregular; teeth not very close-packed; in some specimens tendency to form fourth and fi fth series; unicuspid; normally slightly recurved, occasion- ally almost straight; white or ivory with brown or reddish outer tip; deeply embedded in fl eshy skin tissue (fi g. 7).

Coloration of preserved specimens. (After six months to eighteen years preservation in 75 % ethanol; only slight differences in intensity of coloration discernible between specimens after different periods of conserva- Fig. 6. Close-up view of head of Dicrossus foirni sp. n., holotype tion; fresh material slightly darker, only one specimen (MZUSP 106589), female, showing serrated preoperculum and partially bleached) (fi gs. 1, 2, 6 & 7): Ground colour complete squamation of cheek. of body light brown or (in adult males) pale lead grey; light brownish on forehead, nape, and above lateral line. Scales on body above horizontal line from upper base of pectoral fi n to lower third of caudal-fi n base with brown distal edge, centres in some cases strikingly paler brown, exceptionally ivory. Forehead and infraorbital area brown, nape dark blackish brown. Broad black or grey preorbital stripe, subpreorbital stripe ivory. An- terior section of lips dark grey, lateral sections pale bluish grey, in larger males ivory. Edges of upper and lower jaws grey. Row of four round or oval brown spots extending from upper lateral edge of upper jaw par- allel to anterior edge of cheek to lower edge of eye. Branchiostegal membrane white, cheeks light brown in females and small males, in adult males white to ivory. Preoperculum and operculum light grey with dense dark Fig. 7. Lateral view of typical teeth rows in Dicrossus foirni pigmentation on upper section of preoperculum and sp. n., topotype, (MTD F 32146). adjacent areas. 128 RÖMER et al.: Description of two new cichlid species from Brazil

Tab. 1. Basic statistics of biometric data taken from type specimens of Dicrossus foirni sp. n., and Dicrossus warzeli sp. n. (as % of SL, SL given in mm).

Dicrossus foirni sp. n. Dicrossus warzeli sp. n. Abbreviations n mean min max st.dev n mean min max st.dev SL 7 45.1 26.0 70.9 15.1 9 39.4 28.8 58.8 9.92 standard length HL 7 30.5 28.7 33.0 1.59 9 30,8 27.7 34.0 1.79 head length HD 7 26.5 23.1 32.5 3.27 9 24.2 22.6 25.6 0.84 head depth BD 7 34.1 31.8 37.0 2.26 9 30.2 28.5 31.2 0.96 body depth HW 7 16.7 15.1 17.8 0.88 9 16.0 14.2 16.7 0.83 head width PDL 7 35.4 33.5 38.1 1.54 9 34.0 31.0 36.6 1.49 pre-dorsal length TDL 7 83.5 81.5 85.6 1.48 9 83.5 79.4 87.1 2.32 trans-dorsal length PVL 7 36.9 34.8 41.7 2.72 9 36.1 33.3 39.0 2.07 pre-pelvic length PAL 7 70.8 67.4 75.4 2.73 9 71.9 68.7 76.0 2.53 pre-anal lengt TAL 7 82.6 79.0 85.7 2.23 9 83.6 79.6 86.7 2.67 trans-anal length Eye 7 12.0 10.2 13.9 1.43 9 12.2 10.9 14.4 1.35 eye diameter SNL 7 8.2 5.2 11.0 2.19 9 7.5 5.7 9.1 1.14 snout length CHD 7 6.5 4.8 9.4 1.79 9 5.6 4.5 6.9 0.84 cheek depth POD 7 4.0 3.0 5.5 0.87 9 3.8 3.2 4.8 0.49 pre-orbital depth IOW 7 8.6 7.8 9.7 0.76 9 7.9 7.3 9.0 0.57 inter-orbital width UJL 7 7.5 6.4 8.8 0.99 9 7.1 4.7 8.1 1.04 upper jaw length LJL 7 9.8 8.4 12.1 1.23 9 9.6 6.0 11.8 1.63 lower jaw length CPD 7 13.2 11.8 14.0 0.74 9 12.3 11.3 13.2 0.74 caudal peduncle depth CPL 7 16.4 13.3 19.5 1.92 9 16.2 13.4 18.8 1.63 caudal peduncle length DFB 7 53.7 49.7 57.4 2.48 9 54.2 49.6 58.5 2.47 dorsal fi n base length AFB 7 19.0 16.5 22.3 2.17 9 19.3 16.3 22.6 2.12 anal fi n base length PecL 7 26.0 21.5 28.7 3.11 9 25.2 20.3 27.3 2.27 pectoral fi n length PelL 7 30.6 17.7 56.0 14.30 9 25.4 18.4 35.2 5.01 pelvic fi n length PelSL 7 15.5 13.0 21.7 3.04 9 14.1 12.6 15.7 1.07 pelvic fi n spine length LDS 7 13.3 9.7 20.8 4.27 9 13.4 11.0 15.6 1.38 last dorsal spine length LAS 7 14.3 13.5 15.7 0.82 9 14.0 9.9 16.2 1.99 last anal spine length

Three longitudinal series of round to vertical-ovoid, light brownish or orange brown. Ventral fi ns overall alternating dark grey to blackish, double or triple spots whitish or hyaline, but fi rst membrane with brown or on sides of body; spots grouped on two horizontally black pigmentation. Base of anal fi n whitish, distal parts adjacent scales on centre of lower half of fl ank, on three hyaline blue or translucent, upper soft portion with up adjacent scales midlaterally, and on two or three adja- to eight vertical bars. cent scales about two to three scales below imaginary longitudinal line along middle of dorsum. Seven dorsal Coloration of live specimens (fi gs. 3 – 5, 17 & 18) blotches, one above posterior margin of eye, four along Body of adult males overall pale grey with bluish sheen base of dorsal fi n, and two on caudal peduncle. In some on abdomen. Forehead, interorbital area, and nape grey- larger specimens black spot on dorsum immediately ish or light brownish. Upper abdominal scales with dark below dorsal-fi n rays (6) 7 to 8 (9), extending onto posterior margins. Cheek, lower half of operculum, proximal third to half of dorsal-fi n membranes. Oval throat, and breast ivory. Three longitudinal series of spot on median base of caudal fi n, covering about half double or triple spots on sides. In smaller specimens its height, split horizontally into two equal parts by pale single spots also appear on caudal peduncle. Spots po- grey to whitish stripe on centre. sitioned as described for preserved specimens. Five to All fi ns with irregular pattern of microscopic faint six dorsal blotches along dorsum from nape to base of greyish dots. Dorsal fi n overall milky whitish with faint soft portion of dorsal fi n. Adult dominant specimens submarginal line. Light band running from proximal with variably prominent black spot, occupying up to edge of dorsal-fi n membrane 4 to sub-median position quarter of dorsal-fi n base between spines 6 to 9, which on membranes to membrane 12. Caudal fi n in juveniles not visible in any preserved specimens. Broad preorbital and females overall semi-transparent whitish or hyaline, stripe from orbit to tip of snout. Mature males with row in males upper lobe with vertical stripe pattern distally, of squarish red spots from posterior upper edge of snout i. e. black margin, translucent submarginal band, fol- to lower edge of orbit below posterior edge of pupil. lowed by second black stripe. Scaled base of caudal fi n Upper lip grey in most specimens, but sometimes with Vertebrate Zoology ■ 60 (2) 2010 129 orange or metallic green sheen. Upright oval blotch on stream from the settlement of Sao Luiz (about 4°34′S / 56°15′W), upper edge of preoperculum, in most specimens merg- federal state of Pará, Brazil, preserved after two weeks in the ing into second, half-moon-shaped spot immediately aquarium. posterior on operculum. Paratypes. 8 specimens: MTD F 32152, male, 58.8 mm SL, Mature males with light blue submarginal band along CAS 220 435, female, 33.8 mm SL; MZUSP 106592, female, complete distal edge of dorsal-fi n and upper vertical, 28.8 mm SL; collecting data for all as for holotype, preserved distal half of caudal fi n, submarginal band, in dorsal fi n immediately after importation. MTD F 32154, male, 30.8 mm bordered wine red above and below; round black spots SL, cleared & stained, collecting data as for holotype, preserved in median position on membranes of dorsal spines 7 to after 3 months maintenance in the aquarium. MTD F 32149, 14, second row of spots on proximal part of membranes male, 46.5 mm SL, MTDF 32150, female, 33.4 mm SL, cleared 5 to 15. Caudal fi n in smaller and female specimens and stained, preserved after 5 months maintenance in the aquar- hyaline translucent, in adult males reddish in upper, ium. MTD F 32151, male, 47.7 mm SL, CAS 220 436, female, milky light blue in lower, lobe. Distal part of ventral 30.5 mm SL, collecting data as for holotype, preserved after fi ns white in adult males, hyaline translucent in smaller 7 months in the aquarium. males and in females. During reproduction ventral fi ns of females intense red orange with blue anterior margin. Diagnosis. Dicrossus warzeli sp. n. is a medium-size Anal fi n of mature males bluish with up to eight vertical crenicaratine cichlid species (males up to 60 mm SL, hyaline bars, in females generally hyaline translucent, females to 35 mm SL) sharing all typical characteristics during reproduction with reddish sheen. of the genus Dicrossus. It exhibits clear sexual size and colour dimorphism. The species is unmistakeably Etymology. The species is named in honour of the characterised by three longitudinal rows of dark grey to Federação das Organizações Indígenas do Rio blackish, horizontally elongate, dash-like spots on the Negro. The offi cial abbreviation of the name of this sides of the body; in adult males caudal fi n spatulate. non-govermental organisation is FOIRN. The name ac- knowledges the fact that FOIRN has repeatedly given Description. Morphological data taken from 9 speci- permission to travel on the tribal land of the village mens (28.8 to 58.8 mm SL). For biometric data see communities of different indigenous groups in the mid- tables 1 & 2, for meristic data see table 3. dle and upper Rio Negro and its affl uent rivers, as well as for carrying out observations on wildlife, especially Habitus. (fi gs. 8 – 13 & 19) Body elongate (depth up to insects and fi sh, in these areas. This permitted the ob- about 30 % of SL in adults of both sexes), laterally com- servation and collection of the species described herein. pressed, approximately twice as deep as wide, upright The name of the species is also intended to highlight the oval in frontal view. Predorsal contour regularly arched, fact that the basic human rights of indigenous peoples preventral contour generally straight. Body contour in are still in question in most parts of Amazonia when juvenile males dorsally almost straight, in adult males business projects (such as logging, mining, or the build- slightly convex, ventrally in general slightly concave ing of hydro-electric dams) are planned in the wilder- from ventral fi ns to beginning of anal fi n; in females ness of the Neotropical rainforests. Indigenous peoples body contour dorsally slightly convex, ventrally almost depend on large functional ecosystems that can provide straight, in ripe females slightly convex. Head elongate all necessary resources for indigenous life. For these (head length about 31 % of SL) with small terminal, reasons the founding of FOIRN was a reasonable and narrow, and pointed mouth (as in Dicrossus foirni sp. n., necessary step by the indigenous peoples along the Rio fi g. 5). Snout slightly pointed, comparatively long (about Negro, creating an organisation capable of taking over 7.5 % of SL). Premaxillary fold of lower lip very short. responsibilities and decision-making powers from local Distal edge of preoperculum serrated, with 31 to 42 govermental organisations such as the offi cial Brazilian denticuli (fi g. 12). Cheeks completely scaled. indian agency FUNAI (Fundacao Nacional do Indio). Fins unscaled, except caudal fi n, which has a few scales on its fi rst fi fth to sixth. Dorsal fi n [D. XV.6 (n = 4), XV.7 (n = 3), XV.8 (n = 1), XVII.7 (n = 1)] about half as high as body. Membranes not extending Dicrossus warzeli sp. n. above fi n spines, tips rounded. Dorsal-fi n spines in- creasing rapidly from fi rst to sixth or seventh; seventh (Figs. 8 – 13 & 19, Tabs. 1 – 3) spine about three times longer than fi rst; posterior spines about same length as seventh, increasing in length only slightly to last spine in some specimens. Soft portion of Holotype. (fi g. 8) MZUSP 106591, male 41.2 mm SL, collected dorsal fi n pointed in males, rounded in females. Caudal during September 1992 by F. Warzel. Location.: a small name- fi n [C. 16 (n = 9)] with eight principle rays in each lobe. less right-bank tributary (igarapé) feeding the Rio Tapajós, up- Principle rays increasing in length from upper and lower 130 RÖMER et al.: Description of two new cichlid species from Brazil

Tab. 2. Biometric data taken from Dicrossus foirni sp. n., and Dicrossus warzeli sp. n. (given in mm, for abbreviations see table 1).

Dicrossus foirni sp. n. Sex SL TL HL HD BD HW PDL TDL PPL PAL TAL MZUSP 106589 O HT 42.8 53.4 12.7 10.9 14.3 6.82 14.3 34.9 15.2 26.9 33.8 MZUSP 106590 P PT 57.9 82.3 17.2 16.2 21.4 9.8 20.3 48.5 21.6 36.4 47.9 MTD F 32146 P PT 70.9 97.9 20.3 23.1 26.2 10.7 24.8 58.2 24.8 43.4 57.7 MTD F 32147 O PT 35.0 46.0 10.5 8.7 11.3 6.2 12.8 29.2 12.2 22.3 28.9 MTD F 32148 ? PT 26.0 33.9 8.6 6.1 8.4 4.3 9.9 22.1 10.9 17.8 22.1 CAS 220 433 P PT 45.8 59.0 13.7 12.7 16.2 7.9 15.6 39.2 18.0 30.5 39.2 CAS 220 434 O PT 37.4 48.8 12.1 8.6 11.9 6.4 13.2 31.1 13.1 24.2 30.8 Dicrossus warzeli sp. n. MZUSP 106591 P HT 41.2 53.0 12.5 10.5 11.7 6.8 14.3 32.7 13.7 25.7 32.7 MZUSP 106592 O PT 28.8 36.6 9.0 7.1 8.4 4.7 10.0 24.0 11.0 18.4 24.7 MTD F 32149 P PT 46.5 58.2 13.6 11.1 14.2 7.0 15.7 40.0 16.0 31.4 39.2 MTD F 32150 O PT 33.4 41.5 10.7 8.1 9.9 5.3 11.3 27.5 12.0 21.6 27.2 MTD F 32151 P PT 47.7 65.0 14.2 11.8 14.9 7.7 15.8 41.5 18.5 31.2 41.3 MTD F 32152 P PT 58.8 75.2 16.3 14.1 18.1 8.4 18.2 47.9 20.4 38.1 47.0 MTD F 32154 P PT 33.8 44.3 10.8 7.7 10.4 5.6 12.4 28.7 11.7 23.0 29.0 CAS 220 435 O PT 33.8 42.5 11.5 8.1 10.2 5.7 11.4 28.1 12.4 21.2 28.2 CAS 220 436 O PT 30.5 39.1 9.4 7.6 9.5 5.0 10.5 25.6 11.9 21.1 26.1

Fig. 8. Dicrossus warzeli sp. n., holotype, MZUSP 106591, male, colour pattern in 75 % ethanol, about fi ve years after preservation. margins to centre of fi n. In adult males rays D1 and V1 variably developed (2 to 10 scales), in most cases with slightly less than twice as long as outermost D8 and V8. interruptions, extending on to the base of the caudal fi n. Caudal fi n spatulate in adult males, except rhombic in Teeth in three regular (in all specimens) series in both some exceptionally large specimens; rounded in smaller jaws; teeth close-packed; teeth unicuspid, recurved in males and in females. Anal fi n (A: III.6 (n = 4), III.7 distal third; upper two thirds brown with darker tip, (n = 5) pointed in males, rounded in juveniles and fe- prominent in fl eshy skin tissue (fi g. 13). males. Ventral fi ns [V. I.5 (n = 9)] rounded in juveniles and females, extending back to genital papilla, in larger Coloration of preserved specimens. (described after males pointed with fi lamentous extensions, extending six months to eight years preservation in 75 % etha- back to soft part of anal-fi n base. Pectoral fi ns [P. 11 nol; only slight differences in intensity of coloration (n = 7) 12 (n = 2)] rounded in all specimens examined. discernible between specimens after different periods Scales in E1 row 24 (n = 3) or 25 (n = 5) (one specimen of conservation; fresh material slightly darker) (fi g. 8): lacking scales in E1 row). Upper lateral line (13 to 18 Ground colour medium copper brown; slightly darker scales, one specimen with only 7 scales) continuously above lateral line. Head overall darker than body. Broad arched from head to beginning of soft portion of dorsal black preorbital stripe, subpreorbital stripe light grey. fi n, in some specimens interrupted. Lower lateral line Infraorbital area dark grey, nape dark blackish brown. Vertebrate Zoology ■ 60 (2) 2010 131

Tab. 2. Continuation.

Eye SNL CHD POD IOW UJL LJL CPD CPL DFB AFB PecL Pell PelSL LDS LAS 5.3 2.8 2.4 1.5 3.4 2.8 3.9 5.1 6.4 21.3 7.6 9.4 11.0 9.3 5.4 6.1 6.6 6.2 5.0 3.2 5.6 5.0 6.1 7.9 9.7 32.1 12.9 16.6 26.2 8.6 10.2 9.1 7.2 7.8 6.7 3.4 6.7 6.3 8.6 9.9 11.6 38.9 15.2 20.2 39.7 9.8 14.7 9.7 3.9 1.8 2.1 1.2 2.7 2.3 3.0 4.7 6.1 18.3 6.11 7.5 7.2 4.7 3.4 4.8 3.6 2.0 1.2 0.8 2.3 1.8 2.5 3.6 5.1 14.9 4.3 7.0 6.9 4.4 2.5 3.9 5.2 4.2 2.8 1.9 4.1 3.6 4.1 5.9 6.1 24.4 9.0 13.1 10.4 5.9 5.1 6.5 5.2 2.6 1.9 1.5 3.0 2.8 3.6 4.9 6.2 19.8 6.8 9.8 6.6 5.4 4.2 5.0

4.5 3.7 2.2 1.6 3.7 3.2 3.9 4.8 6.8 20.4 7.5 10.9 12.3 5.4 5.8 4.1 3.6 1.6 1.4 1.2 2.1 1.8 3.1 3.6 5.0 15.6 5.3 5.9 6.6 3.9 3.9 4.2 5.3 3.5 3.2 2.2 3.5 3.4 4.6 6.1 7.6 26.1 10.5 10.8 8.5 5.9 5.1 5.8 3.8 2.1 1.7 1.1 2.5 2.5 3.6 4.0 6.3 17.5 5.7 8.3 8.4 4.9 4.1 5.4 5.2 4.1 2.8 1.9 3.8 3.6 4.5 6.3 6.4 27.9 10.5 12.4 16.8 7.5 7.1 7.3 6.7 4.6 4.1 2.3 4.3 4.8 5.2 6.7 9.8 31.8 12.1 14.8 12.1 7.9 9.2 7.8 4.8 2.3 1.8 1.2 2.9 2.6 3.3 4.4 5.2 18.8 6.4 9.1 9.0 4.7 4.5 5.0 4.9 2.5 1.5 1.1 2.8 1.6 2.0 3.8 4.8 18.1 5.5 9.2 8.9 4.7 4.3 4.5 4.0 2.7 1.7 1.1 2.3 2.2 3.6 3.6 5.3 16.6 6.1 8.3 7.3 4.7 4.2 4.9

Fig. 9. Dicrossus warzeli sp. n., adult male, dominant, territorial, not preserved. Photo: Frank Warzel.

Anterior section of lips dark grey, lateral sections pale most cases three longitudinally adjacent scales in row. grey to whitish, edges of upper and lower jaws grey. Intervals between dash-like spots about equal, except Branchiostegal membrane grey, cheek light brown, pre- often narrower on caudal peduncle. Vertical intervals operculum and operculum grey with dense pigmentation between rows of dash-like spots about one scale wide. composed of numerous black dots on preoperculum and Series of seven dorsal spots, one above posterior margin adjacent areas. of eye, four along base of dorsal fi n often merging into Three longitudinal series of horizontal, dash-like fi rst series of dash-like spots below it, two on caudal spots on sides, fi rst extending in straight line from dor- peduncle, not completely visible in all specimens, in sal edge of orbit to upper scales of caudal peduncle, some specimens adjacent spots may be merged. Black second from upper edge of operculum onto imaginary spot of variable shape, mostly oval, on dorsum imme- line between upper and middle thirds of caudal base, diately below and on lower fi fth of membranes between third from behind upper edge of pectoral fi n insertion dorsal-fi n rays 7 and 8, most prominent where coinci- onto imaginary line between middle and lower thirds of dent with greyish lateral band on E1 scales, extending caudal base, both second and third terminating on fi rst from upper edge of operculum to distal edge of caudal quarter of caudal fi n. Individual dash-like spots forming peduncle. Squarish spot on centre of caudal-fi n base, lateral series, each covering centres of two to four, in covering about half depth of latter height. 132 RÖMER et al.: Description of two new cichlid species from Brazil

Fig. 10. Dicrossus warzeli sp. n., juvenile female, subdominant, showing typical clear dash line pattern, not preserved. Photo: Frank Warzel.

Fig. 11. Dicrossus warzeli sp. n., adult female, dominant, brood care coloration, guarding fry, not preserved. Photo: Frank Warzel.

All fi ns with irregular pattern of microscopic Coloration of live specimens. (fi gs. 9 – 11 & 19) black dots. Dorsal fi n overall milky whitish with Body of adult males overall beige; upper half of head light submarginal stripe; central parts of membranes and abdomen greyish or light brown, partially light along hard rays pigmented dusky grey in females, in mossy green in aggression; ventral region, cheeks, males bordered translucent milky white; males with and throat ivory. Three longitudinal series of dash-like light submarginal stripe along entire fi n, continuing spots on sides (fi g. 10 & 11), positioned as described on caudal fi n. Caudal fi n in males with faint sub- for preserved specimens. Four dorsal spots along marginal stripe (rather diffi cult to detect) on distal edge dorsal-fi n base. Adult dominant or aggressive males of upper lobe; remainder of caudal fi n overall semi- with prominent black spot covering up to a fi fth of the transparent hyaline, rarely milky white; scaled base height of dorsal fi n between spines 6 to 8, also present in of caudal fi n pale yellowish brown (females) or ash females guarding fry. Broad preorbital stripe from grey (most males). Ventral fi ns overall hyaline whit- orbit to tip of snout, in some female specimens extend- ish, with pattern of small pigmented spots along soft ing onto upper surface of anterior half of snout. Upper rays, more concentrated along spine. Base of anal fi n lip metallic blue to shiny turquoise in mature males, whitish in males, hyaline translucent in females, upper greyish to pale orange in females. Upright oval blotch soft portion with up to four vertical bars. on upper edge of preoperculum, second triangular to Vertebrate Zoology ■ 60 (2) 2010 133 n fi n n n n (s): PF: (h): soft rays hard rays pelvic fi

Fig. 12. Close-up view of head of Dicrossus warzeli sp. n., para- type (MTD F 32149), male, demonstrating serrated preoperculum and complete squamation of cheek (cleared & stained) 33.830.5 15 15 6 6 3 3 6 6 1 1 5 5 12 11 16 16 24 25 37.4 15 7 3 7 1 5 12 16 25 45.8 16 6 3 7 1 5 11 16 24 33.8 17 7 3 7 1 5 11 16 25 35.0 15 7 3 7 1 5 11 16 24 AF: 47.758.8 15 15 7 7 3 3 6 7 1 1 5 5 11 11 16 16 24 24 33.4 15 6 3 6 1 5 11 16 25 caudal fi 70.9 15 7 3 7 1 5 11 16 26 dorsal fi 46.5 15 8 3 7 1 5 12 16 25 CF: 57.9 15 728.8 3 15 6 6 1 3 5 7 11 1 16 5 26 11 16 DF : def. pectoral 42.8 15 741.2 3 15 7 7 1 3 5 7 11 1 16 5 25 11 16 25 PecF: O PT O PT O PT P PT P PT O PT P PT P PT O PT P PT P PT P PT O PT O HT P HT sp. n. (SL given in mm, for abbreviations see last column of table). warzeli sp. n. (SL foirni sp. n., and Dicrossus Dicrossus

Fig. 13. Lateral view of typical teeth rows in a male Dicrossus warzeli sp. n. paratype (MTD F 32149). CAS 220435 CAS 220436 CAS 220434 CAS 220433 MTD F 32148 ? PT 26.0 15 7 3 7 1 5 11 16 25 anal fi MTD F 32154 MTD F 32147 MTD F 32151 MTD F 32152 MTD F 32150 MTD F 32146 MTD F 32149 MZUSP MZUSP MZUSP Collection NumberMZUSP Sex SL DF (h) DF (s) MZUSP AF (h) AF (s) PF (h) PF (s) PecFhalf-moon-shaped CF LRS spot Abbreviations immediately posterior on oper-

culum.

sp. n. sp.

sp. n. sp. Aggressive specimens with prominent brown to

warzeli Dicrossus Dicrossus foirni Dicrossus black, rarely mossy green spot on operculum, usually

Tab. 3 . Meristic data taken from Tab. occurring in conjunction with dark band on proximal 134 RÖMER et al.: Description of two new cichlid species from Brazil

Fig. 14. Map of Brazil showing distribution of Dicrossus foirni sp. n. (type locality: white star in red dot; other collecting sites: stars in white dots), and Dicrossus warzeli sp. n.. (type region: red square; other collecting sites: black dots in white squares). part of dorsal fi n. Also in aggression, conspicuous black his numerous fi eld trips to Brazil and Colombia. In spot on dorsum below and on lower fi fth of dorsal-fi n his later years he also focussed on dwarf cichlids of membranes 6 to 9, usually when greyish midlateral band the genera Teleocichla, , and Dicros- also apparent; some specimens also exhibiting faint sus. Born on December 21st 1960, Frank unexpect- round black lateral spot directly behind gill cover, and edly passed away, much too early, after heart surgery another below dorsal-fi n spines 8 and 9. Light blue to on April 27th 2004. metallic green sub-marginal band along entire distal edge of dorsal fi n and upper distal half of caudal fi n. Caudal fi n overall hyaline translucent, in adult males milky white, in some cases with reddish base and blu- Geographical distribution and ecological ish distal portion. Distal part of ventral fi ns in adult notes males bluish, in females hyaline translucent; during reproduction, in males overall light violet or reddish with turquoise anterior margin, in females yellowish to As far as is known at present, Dicrossus foirni sp. n. intense orange. Anal fi n of mature males bluish violet occurs in the middle to upper Rio Negro system (fi g. to wine red, in females generally hyaline translucent. 14). Collecting sites are restricted exclusively to left- hand tributaries of the main channel of the Rio Negro, Etymology. The species is named in honour and from the mouth of the Rio Branco in the east upriver to memory of the skilled German aquarist Frank Mar- the settlement of Santa Isabell in the west. Observation tin Warzel. Frank not only collected the type speci- or collecting has been reported from the drainages of mens, but was also the fi rst to import the species from the Rios Ararira, Demini, Marauiá, Padauiri, and Preto the Rio Tapajos to Germany, as well as to observe its (Kullander, 1984, 1990; Rank, 1994; own observa- behaviour, including reproduction, in the fi eld and in tions). the aquarium (Warzel, 1996). He dedicated most of Dicrossus foirni sp. n. has been found in all types his life to research on Neotropical cichlids, especially of water. Juveniles have been found exclusively in very those of the genus Crenicichla. Dozens of publica- soft and acid, clear and black water (Windisch, 1992; tions dealing with Neotropical cichlids resulted from Wilhelm, pers. comm. 2007, 2008; own observations), Vertebrate Zoology ■ 60 (2) 2010 135

while sub-adults have also been found in whitewater brooks (Eon, pers. comm., May 2010). In 1992 UR observed half-grown individuals shoaling with several hundred Dicrossus fi lamentosus in the mouth of the Igarapé Prósperitáte (type locality), lower Rio Preto, and in 1994 adult specimens in only small numbers in a shallow forest lake about 30 minutes away by canoe from that location (fi g. 15). In the Igarapé Prósper- itáte Dicrossus foirni sp. n. were found together with the cichlids Aequidens sp., Apistogramma gibbiceps Meinken, 1969, Apistogramma paucisquamis Kul- lander & Staeck, 1988, Apistogramma pertensis (Haseman, 1911), Crenicichla inpa Ploeg, 1991, Cre- ni cichla notophthalmus Regan, 1913, the characins Ho- plias malabaricus (Bloch, 1794), and Paracheirodon axelrodi (Schultz, 1956). More detailed data on habitat structure, sociobiology, and further biotic and abiotic factors may be found in Römer (1992a & b, 1994, 1998, 2000). Dicrossus warzeli sp. n. seems to be endemic to the middle to upper Rio Tapajos system (Kullander, 1984, Warzel, 1995, Stawikowski & Werner, 2004) (fi g. 14). Collecting sites are located in small igarapés near São Luiz (type locality) (Warzel, in lit.) (fi g. 16), in the Igarapé Pimental (Kullander, 1990; Warzel, in Fig. 15. View of typical habitat of Dicrossus foirni sp. n., Station lit.), and in a small river about 56 km south of Baburé F1/91R, lower Igarapé Prósperitáte, March 1994. (Stawikowski & Werner, 2004). Dicrossus warzeli sp. n. has been found exclusively in clearwater rivers. Frank Warzel collected several juveniles of about 20 mm SL in small brooks with sandy bottoms and patches of leaf litter. The fi sh were search- ing the bottom substrate for small invertebrates and detritus by picking up and turning the leaves. These fi sh have been collected exclusively in very soft water (no detectable hardness) with high acidity (pH value regularly measured at less than 5, in most cases less than 4). Intensive fi eld research on the ecology of both species is still required. There is no evidence of any overlap of the ranges of the two species. Remarkably, Dicrossus warzeli sp. n. and Dicrossus maculatus seem to be the only species of the genus known exclusively from south of the main channel of the Amazon river. Compared to the species described here, Dicrossus maculatus is known to be far less sensitive to shifting water conditions, especially pH and water hardness. Dicrossus warzeli sp. n., by contrast, seems to be stenoecious (restricted to very low acidity values and extreme soft water). Initially this also seemed to be the case with Dicrossus foirni sp. n., but surprisingly more recent observations of the species (Eon, pers. comm. May 2010) contradict this earlier assumption (cf. Römer, 2000; Staeck 1993). Fig. 16. View of typical habitat of Dicrossus warzeli sp. n., small tributary of lower Tapajós upriver from cachoeiras (rapids) near Sao Luiz, September 1992. Photo: Frank Warzel. 136 RÖMER et al.: Description of two new cichlid species from Brazil

Fig. 17. Dicrossus foirni sp. n., juvenile male, dominant, showing colour pattern of “bleeding” spots, not preserved.

Fig. 18. Dicrossus foirni sp. n., juvenile male, aggressive pattern, not preserved.

Discussion other cichlid species known to date can be mistaken for either of the two. Sharing several morphological and behavioural traits, Dicrossus foirni sp. n. and Dicrossus The genus Dicrossus consists of at least fi ve distinct warzeli sp. n. appear to be more closely related to each species, which are unmistakable amongst all other Neo- other than to other members of the genus. Detailed ge- tropical cichlid genera and species. Dicrossus foirni netic studies of the phylogenetic relationships between sp. n. and Dicrossus warzeli sp. n. are the fourth and the species of the genus are in progress. fi fth species described. The genus Dicrossus apparently Live as well as preserved specimens of Dicros- falls into two major groups: on the one hand Dicros- sus foirni sp. n. and Dicrossus warzeli sp. n. can be sus fi lamentosus, Dicrossus gladicauda, and Dicrossus easily distinguished from one another by their clearly maculatus, with two rows of large spots in each case, different spot patterns. Dicrossus foirni sp. n. exhibits while on the other hand Dicrossus foirni sp. n. and Di- longitudinal rows of upright oval, double spots versus crossus warzeli sp. n. exhibit three rows of, respectively, horizontal dash-like spots in Dicrossus warzeli sp. n.. double spots or dash-like spots on the sides of the body. Furthermore, in smaller aggressive or adult dominant The colour patterns of the two species described here- males of Dicrossus foirni sp. n. the spots in the two in are unique amongst Neotropical cichlids, hence no lower rows appear to be “bleeding” vertically (fi g. 17) Vertebrate Zoology ■ 60 (2) 2010 137

Fig. 19. Dicrossus warzeli sp. n., adult male, aggressive pattern, not preserved. Note the narrow and pointed mouth of this species, visible here. Photo: Frank Warzel.

38 sp. n. exhibits a distinct lateral band combined with a prominent spot on the operculum and a dark stripe on

36 the proximal part of the dorsal fi n (fi g. 19). Statistical analysis of biometric data taken from pre- served specimens shows that both species are similar 34 overall. However, relative body depth plotted against relative head width reveals signifi cantly differentiated 32 polygons for Dicrossus foirni sp. n. and Dicrossus warzeli sp. n. (fi g. 20). Tooth morphology is another 30 diagnostic character, especially useful in small preserved specimens. 28 With the description of Dicrossus foirni sp. n. and Dicrossus warzeli sp. n. all taxonomic problems in the 14 15 16 17 genus Dicrossus might appear to be resolved. How- Fig. 20. Scatterplot of biometric data for Dicrossus foirni sp. n. ever, when Kullander (1978) re-described Dicros- (circles), and Dicrossus warzeli sp. n.. (stars), (x-axis: head width, sus fi lamentosus he used material exclusively from the y-axis: body depth, both given as % of SL). Missing overlap of Colombian and Venezuelan Rio Orinoco drainage. But polygons represents diagnostic stage of data sets given in tabs. the original type material of the species originates from 1 & 2. the Brazilian Rio Negro. In fact Dicrossus fi lamentosus from the middle and upper Rio Negro system are easily distinguished from specimens of the Orinoco population. There are at least signifi cant differences in the pattern of black lateral and merging into one another via more or less intense blotches and in the coloration of the caudal fi n. In ad- grey to blackish vertical stripes, which often have a dition, specimens from the Rio Negro seem to be more washed-out appearance. Differences in the aggressive slender than those collected from the Colombian and colour patterns of the males of the two species also Venezuelan Rio Orinoco drainage. More importantly, offer diagnostic features. While the aggressive pattern preliminary genetic studies (Ready, in lit.) have also of Dicrossus foirni sp. n. is dominated by vertical bars indicated differences between specimens from the two which often look washed out (fi g. 18), Dicrossus warzeli regions. 138 RÖMER et al.: Description of two new cichlid species from Brazil

The material available for study is still too limited to Mayland, H.J. & Bork, D. (1997): Zwergbuntbarsche: Süd- produce suffi cient data to confi rm the status of the two amerikanische Geophaginen und Crenicarinen. – Land- populations, but there does seem to be a strong prob- buch Verlag, Hannover, Germany:164 – 171. ability that the specimens on which the description and Römer, U. (1992a): Weitere Beobachtungen zur Verbrei tung, re-description are based represent two different species. Ökologie und Aquarienbiologie von Lae ta cara spec. With this in mind, the two populations should be treated „Orangefl ossen”. – DCG-Informationen, 23(4): 74 – 83. as potentially different species: Dicrossus fi lamentosus Römer, U. (1992b): Weitere Beobachtungen zu Verbreitung from the Brazilian Amazon system and Dicrossus sp. “C” und Ökologie des „Orangefl ossen”-Laetacara im Rio from the Orinoco drainage, awaiting description. Further Negro-Gebiet (NW-Brasilien). – DCG-Informationen, research to resolve this dilemma is strongly encouraged. 23(12): 245 – 255. Römer, U. (1994a): Laetacara spec. „Orangefl ossen”: Er- gän zende Beobachtungen zur Verbreitung und Ökologie im Gebiet des Rio Negro (NW-Brasilien). – DCG-Infor- Acknowledgements mationen, 25(12): 274 – 288. Römer, U. (1994b): Einige Aspekte der Mikrohabitatwahl südamerikanischer Zwergbuntbarsche (Teleostei: Cich- We are thankful to Manfred Geismann (Cologne, FRG), lidae). Pp. 239 – 254. In: Greven, H. & Riehl, R. Stefan Leisner (Ulm, FRG), Axel Schneider (Offenburg, (eds.): Verhalten der Aquarienfi sche. – Düsseldorf. Bir- FRG), Frank Warzel (†), and Martin Wöhler (Bielefeld, git Schmettkamp Verlag, Bornheim, Germany. FRG) for collecting and providing preserved and live material Römer, U. (2000): Cichlid Atlas, Vol. 1: Natural History for this study. Horst Warzel kindly transferred the complete of South American Dwarf Cichlids. – Mergus, Melle, estate of his son Frank to the senior author, permitting the Germany, 1311 pp. illustration of live specimens of Dicrossus warzeli sp. n.. The Römer, U. (2006): Cichlid Atlas, Vol. 2: Natural History of Federação das Organizações Indígenas do Rio Negro South American Dwarf Cichlids, Part 2. – Mergus, Mel- (FOIRN) (Sao Gabriel da Cachoeira, Brazil) kindly gave per- le, Germany, 1319 pp. mission to travel in the tribal lands of the indigenous peoples Römer, U. & Hahn, I. (2008): Apistogramma barlowi sp. n.: of the middle and upper Rio Negro, as well as for observ- Description of a new facultative mouth-breeding cich- ing and collecting dwarf cichlids in that area. Dr. Wolfgang lid species (Teleostei: Perciformes: Geophaginae) from Beisenherz (Bielefeld University, FRG) kindly provided Northern Peru. – Vertebrate Zoology, 58(1): 49 – 6. working facilities for behavioural studies. We also thank Mary Römer, U., Hahn, I., Melgar, J., Soares, D.P., & Wöh- Bailey (Crediton, Devon, UK) for reading and commenting ler, M. (2004): Redescription of Apistogramma erem- on an earlier version of this manuscript. nopyge Ready & Kullander, 2004. – Das Aquarium, 38(12) (426): 17 – 34. Römer, U., Hahn, I., Römer, E., Soares, D.P. & Wöhler, M. (2003): Apistogramma martini sp. n. – Beschreibung References eines geophaginen Zwergcichliden (Teleostei: Perci for- mes) aus dem nördlichen Peru. – Das Aquarium, 37(4) (406): 14 – 29. Kullander, S.O. (1978): A redescription of Crenicara fi - Schindler, I. & Staeck, W. (2008): Dicrossus gladicauda la mentosa Ladiges, 1958 (Teleostei: Cichlidae). – Mitt. sp. n. – a new species of crenicarine dwarf cichlids (Te- Hamb. Zool. Mus. Inst., 75: 267 – 278. leo stei: Perciformes: Cichlidae) from Colombia, South- Kullander, S.O. (1986): Cichlid fi shes of the Amazon America. – Vertebrate Zoology, 58(1): 67 – 73. River drainage of Peru. – Swedish Museum of Natural Staeck, W. (2003): Cichliden Lexikon, Teil 3: Süd ameri- History, Stockholm, 431 pp. kanische Zwergbuntbarsche. – Dähne Verlag, Ettlingen, Kullander, S.O. (1990): Mazarunia mazarunii (Teleostei: Germany: 139 – 142. Cichlidae), a new genus and species from Guyana, South Stawikowski, R. & Werner, U. (2004): Die Buntbarsche America. – Ichthyol. Expl. Freshwaters, 1(1): 3 – 14. Amerikas, Band 3: Erdfresser, Hecht- und Kammbunt- Ladiges, W. (1958a): Bemerkungen zu einigen Neu impor- barsche. – Verlag Eugen Ulmer, Stuttgart, Germany: ten. – Die Aquarien- und Terrarienzeitschrift (DATZ), 54 – 71. 11(7): 203 – 204. Tomey, W.A. (1983): Een onbekende Crenicara? – Het A- Ladiges, W. (1958b): Crenicara fi lamentosa spec. nov. – qua rium, 53(7/8): 174 – 178. ein neuer seltener Cichlide aus Südamerika. – Int. Rev. Warzel, F. (1996): Ein neuer Schachbrettcichlide aus dem Hydro biol., 44(2): 299 – 302. Tapajós. – Das Cichlidenjahrbuch 6 (1996): 80 – 82. Linke, H. & Staeck, W. (1992): Amerikanische Cichliden Windisch, W. (1992): Dicrossus sp. “Rio Negro”. – Die A- I – Kleine Buntbarsche. – Tetra-Verlag, Melle, Germa- quarien- und Terrarienzeitschrift (DATZ), 45(12): 767 – ny, 232 pp. 770.